Integrative Musterbildung kranialer Strukturen

Form und Gestalt kraniofazialer Strukturen sind primär beeinflusst durch die inhärente Integration der unterschiedlichsten Funktionssysteme und externer selektiver Einflüsse. Die Variabilität der Schädel-Morphe ist ein Indikator für solche Einflussfaktoren und damit ein idealer Gegenstand für vergleichende Analysen morphogenetischer Formbildung. Zur Ermittlung morphologisch-adaptiver Trends und Muster wurden sowohl Hypothesen zur morphologischen Differenziertheit als auch zu Korrelationen zwischen modularen Schädelkompartimenten (fazial, neurokranial, basikranial) untersucht. Zusätzlich wurden aus Schichtröntgenaufnahmen (CT) virtuelle Modelle rekonstruiert, welche die Interpretation der statistischen Befunde unterstützen sollten. Zur Berechnung der Gestaltunterschiede wurden mittels eines mechanischen Gelenkarm-Messgerätes (MicroScribe-G2) max. 85 ektokraniale Messpunkte (Landmarks) bzw. dreidimensionale Koordinaten an ca. 520 Schädeln von fünf rezenten Gattungen der Überfamilie Hominoidea (Hylobates, Pongo, Gorilla, Pan und Homo) akquiriert. Aus dem Datensatz wurden geometrische Störfaktoren (Größe, Translation, Rotation) mathematisch eliminiert und die verbleibenden Residuale bzw. ‚Gestalt-Variablen‘ diversen multivariat-statistischen Verfahren unterzogen (Faktoren, Cluster-, Regressions- und Korrelationsanalysen sowie statistische Tests). Die angewandten Methoden erhalten die geometrische Information der Untersuchungsobjekte über alle Analyseschritte hinweg und werden unter der Bezeichnung „Geometric Morphometrics (GMM)“ als aktueller Ansatz der Morphometrie zusammengefasst. Für die unterschiedlichen Fragestellungen wurden spezifische Datensätze generiert. Es konnten diverse morphologische Trends und adaptive Muster mit Hilfe der Synthese statistischer Methoden und computer-basierter Rekonstruktionen aus den generierten Datensätzen ermittelt werden. Außerdem war es möglich, präzise zu rekonstruieren, welche kranialen Strukturen innerhalb der Stichprobe miteinander wechselwirken, einzigartige Variabilitäten repräsentieren oder eher homogen gestaltet sind. Die vorliegenden Befunde lassen erkennen, dass Fazial- und Neurokranium am stärksten miteinander korrelieren, während das Basikranium geringe Abhängigkeiten in Bezug auf Gesichts- oder Hirnschädelveränderungen zeigte. Das Basikranium erweist sich zudem bei den nicht-menschlichen Hominoidea und über alle Analysen hinweg als konservative und evolutiv-persistente Struktur mit dem geringsten Veränderungs-Potential. Juvenile Individuen zeigen eine hohe Affinität zueinander und zu Formen mit einem kleinem Gesichts- und großem Hirnschädel. Während das Kranium des rezenten Menschen primär von Enkephalisation und fazialer Retraktion (Orthognathisierung) dominiert ist und somit eine einzigartige Gestalt aufweist, zeigt sich der Kauapparat als maßgeblich formbildendes Kompartiment bei den nicht-menschlichen Formen. Die Verbindung von GMM mit den interaktiven Möglichkeiten computergenerierter Modelle erwies sich als valides Werkzeug zur Erfassung der aufgeworfenen Fragestellungen. Die Interpretation der Befunde ist durch massive Interkorrelationen der untersuchten Strukturen und der statistisch-mathematischen Prozeduren als hoch komplex zu kennzeichnen. Die Studie präsentiert einen innovativen Ansatz der modernen Morphometrie, welcher für zukünftige Untersuchungen im Bereich der kraniofazialen Gestaltanalyse ausgebaut werden könnte. Dabei verspricht die Verknüpfung mit ‚klassischen’ und modernen Zugängen (z. B. Molekularbiologie) gesteigerte Erkenntnismöglichkeiten für künftige morphometrische Fragestellungen.

L’orientation de la cavité glénoïde de la scapula chez les grands singes (Gorilla, Pan et Pongo) et l’humain moderne : une étude comparative et ontogénique

Le degré de rétention de l’arboricolisme dans le répertoire locomoteur des hominines fossiles du Pliocène est toujours matière à débat, les études ayant principalement porté sur la courbure des phalanges et la proportion des membres. Vu la récente découverte de DIK-1-1 (A. afarensis) et de la scapula qui lui est associée, l’étude de cet os d’un point de vue fonctionnel est intéressante, puisqu’il est directement impliqué dans la locomotion de presque tous les hominoïdes. Le but de cette étude est de tenter d’établir un lien entre l’orientation supéro-inférieure (SI) et antéro-postérieure (AP) de la cavité glénoïde de la scapula et les comportements locomoteurs chez les grands singes et l’humain moderne. Des analyses comparatives sur les adultes ont été réalisées pour 1) voir s’il existe des différences dans la morphologie étudiée entre les espèces et 2) voir si ces différences peuvent être expliquées par la taille corporelle. Des analyses ontogéniques ont aussi été réalisées pour voir si un accroissement de la taille corporelle pendant le développement et les changements locomoteurs qui y sont associés correspondent à un changement d’orientation de la cavité glénoïde. Les résultats montrent que les humains ont une cavité glénoïde qui est orientée moins supérieurement que les grands singes, mais que Pongo, bien qu’étant le plus arboricole, n’a pas l’orientation la plus supérieure. Les « knuckle-walkers » (Pan et Gorilla) se distinguent des autres hominoïdes avec une orientation de la surface glénoïde relative à l’épine plus inférieure. La taille corporelle ne semble pas influencer la morphologie étudiée, sauf parfois chez le gorille. Seuls l’humain et les mâles Pongo montrent un changement ontogénique dans l’orientation de la cavité glénoïde relativement à l’épine. Sur la base de ces résultats, l’orientation de la cavité glénoïde semble refléter partiellement la fonction du membre supérieur dans la locomotion, mais des recherches plus poussées sont nécessaires. Mots-Clés : Scapula, cavité glénoïde, grands singes, humains, locomotion, arboricolisme.

Inferences regarding the diet of extinct hominins: structural and functional trends in dental and mandibular morphology within the hominin clade

This contribution investigates the evolution of diet in the Pan – Homo and hominin clades. It does this by focusing on 12 variables (nine dental and three mandibular) for which data are available about extant chimpanzees, modern humans and most extinct hominins. Previous analyses of this type have approached the interpretation of dental and gnathic function by focusing on the identification of the food consumed (i.e. fruits, leaves, etc.) rather than on the physical properties (i.e. hardness, toughness, etc.) of those foods, and they have not specifically addressed the role that the physical properties of foods play in determining dental adaptations. We take the available evidence for the 12 variables, and set out what the expression of each of those variables is in extant chimpanzees, the earliest hominins, archaic hominins, megadont archaic hominins, and an inclusive grouping made up of transitional hominins and pre-modern Homo . We then present hypotheses about what the states of these variables would be in the last common ancestor of the Pan – Homo clade and in the stem hominin. We review the physical properties of food and suggest how these physical properties can be used to investigate the functional morphology of the dentition. We show what aspects of anterior tooth morphology are critical for food preparation (e.g. peeling fruit) prior to its ingestion, which features of the postcanine dentition (e.g. overall and relative size of the crowns) are related to the reduction in the particle size of food, and how information about the macrostructure (e.g. enamel thickness) and microstructure (e.g. extent and location of enamel prism decussation) of the enamel cap might be used to make predictions about the types of foods consumed by extinct hominins. Specifically, we show how thick enamel can protect against the generation and propagation of cracks in the enamel that begin at the enamel– dentine junction and move towards the outer enamel surface.

Influence de la locomotion sur la morphologie de l’articulation distale de l’humérus chez les hominoïdes

La masse corporelle et la direction des charges sont des facteurs qui peuvent modifier la morphologie des surfaces articulaires qui sont généralement orientées et de taille suffisante pour résister aux charges chroniques. Chez les hominoïdes, les forces de tension et compression, générées par la locomotion, sont transmises à travers l’articulation du coude. Ces espèces ont une morphologie similaire de l’extrémité distale de l’humérus, mais qui présente certaines différences selon la taille des individus et leurs modes de locomotion. Ce projet tente de caractériser plus exhaustivement cette variation en analysant la largeur des surfaces articulaires ainsi que leur position et orientation par rapport à l’axe long de la diaphyse. La prémisse de ce mémoire est que, chez les espèces plus arboricoles, la morphologie de l’articulation distale de l’humérus répond aux stress transverses générés par les puissants muscles fléchisseurs du poignet et des doigts qui traversent le coude obliquement. En revanche, les espèces plus terrestres présentent une morphologie permettant de résister aux forces axiales provenant du contact avec le sol. Des coordonnées tridimensionnelles et des mesures linéaires ont été recueillies sur un échantillon squelettique d’individus des genres Homo, Pan, Gorilla et Pongo. Les résultats obtenus révèlent que l’orientation et la position des surfaces articulaires de la trochlée correspondent aux types de locomotion, or leur taille et celle et du capitulum semblent être influencées par la taille des individus. L’hypothèse suggérant que les stress reliés aux divers modes de locomotion des hominoïdes influencent la morphologie de l’articulation distale de l’humérus est donc supportée.

Do chimpanzees (Pan troglodytes) and 2-year-old children (Homo sapiens) understand double invisible displacement?

Chimpanzees (Pan troglodytes) and young children (Homo sapiens) have difficulty with double invisible displacements in which an object is hidden in two nonadjacent boxes in a linear array. Experiment 1 eliminated the possibility that chimpanzees' previous poor performance was due to the hiding direction of the displacement device. As in Call (2001), subjects failed double nonadjacent displacements, showing a tendency to select adjacent boxes. In Experiments 2 and 3, chimpanzees and 24-month-old children were tested on a new adaptation of the task in which four hiding boxes were presented in a diamond-shaped array on a vertical plane. Both species performed above chance on double invisible displacements using this format, suggesting that previous poor performance was due to a response bias or inhibition problem rather than a fundamental limitation in representational capacity.

Structure analysis of CCR5 from human and primates

It is well known that the chemokine receptor CCR5 plays very important roles in HIV-1 virus infection. A three-dimensional molecular model of human CCR5 was generated by SYBYL, a distance geometry-based homologous modeling package, using the corresponding transmembrane domain of bacteriorhodopsin as the template. On the basis of human CCR5 model, we also built 18 3D molecular models of CCR5 in primates from Pongo pygmaeus, Pygathrix nemaeus, Macaca assameniss, Trachy-pithecus phayrei, T. francoisi, M. arotoides, Rhinopithecus roxellance, R, bieti, R. avunculus, Hylobates leucogenys, Pan troglodytes, Gorilla gorilla, Cercopithecus aethiops 1, C. aethiops 2, Papio hamadryas M. mulatta, M. fascicularis and M. nemestrina. Structural analyses and statistics results suggested that the main-chains of the primate CCR5 were similar to that of the human CCR5 and that the fit-RMS deviation values of these primate CCR5 were less than 0.1 Angstrom. Moreover, the structures of these CCR5 proteins, except those of the African green monkey 1 (C.aet1), do not have a remarkable difference. It is proved that the 14th residue is possibly very important in the inhibition infections by M-tropic HIV-1, and it is also demonstrated that the 13th residue of human CCR5 was changed from asparagine into aspartic acid in all these primates. It means that the primate CCR5 no longer depend on CD4 for efficient entry, but human CCR5 may have evolved subsequently due to the use of CD4 as a receptor, allowing the high-affinity chemokine receptor-binding site of HIV to be sequestered from host immune surveillance. (C) 2000 Elsevier Science B.V. All rights reserved.

猕猴白化病分子机制和灵长类的分子进化研究

酪氨酸酶是黑色素合成当中的关键酶。人酪氨酸酶基因包括5个外显子，在染色体11q14-q21位置上占据了约50kb长的区域。对人类眼皮肤型白化病（Oculocutaneous albinism, OCA)的许多研究表明，该病主要是由于酪氨酸酶基因的突变引起的。昆明动物研究所白化猴研究小组数十年来一直从事白化猕猴的培育和研究工作，目前饲养着2只白化猕猴和它们的后代，这提供了我们研究猕猴白化分子机制目的条件。为了弄清猕猴白化病的分子机制，我们根据人酪氨酸酶基因序列设计了5对PCR引物扩增相应的5个外显子，序列分析表明，白化猕猴珍珍酪氨酸酶基因第184个密码子第2位置（外显子1的核苷酸位置551）处发生一个C→A的无义突变，使编码丝氨酸（Ser)的密码子变成了一个终止密码，这样后面1038bp的核苷酸片段（346个氨基酸残基）被截断，导致酪氨酸酶翻译不完全，迄今为止，并没有发现合成黑色素的第二条生化途径，因此由于该酶不能行使正常功能而将导致黑色素不能正常表达。这可能是导致该例猕猴白化病的原因。为了解酪氨酸酶基因序列变异的规律及其与功能的关系，探讨该基因作为系统发育研究中遗传标记的有效性，我们测定了黑猩猩（Pan troglodytes)、倭黑猩猩（Pan paniscus)、大猩猩（Gorilla gorilla)、猩猩(Pongo pygmaeus)、长臂猿（Hylobates lar)、食蟹猴（Macaca fascicularis)、狒狒（Simia cynocephalus)、猕猴（Macaca mulatta)、熊猴（Macaca assamensis)、菲氏叶猴（Presbytis p. crepusculus)、白臀叶猴（Pygathrix nemaeus)、滇金丝猴（Rhinopithecus r. bieti）和蛛猴（Ateles paniscus)13个灵长类中代表种的酪氨酸酶基因全部5个外显子的DNA序列。基于这些序列，用简约法构建了分子系统树。结果表明，人猿超科与旧大陆猴各自形成一单系群。人猿超科各物种和旧大陆猴有明显分化，人与大猩猩的关系比人与黑猩猩的关系近。酪氨酸酶基因在解决灵长类系统发育关系上是一个较有用的基因。为了进一步了解中国猕猴（Macaca mulatta)的亚种分化和不同地理群体间的基因流状况，我们测定了来自中云南、广西、福建、海南、浙江、河南、湖南、湖北、安徽、四川、贵州和越南猕猴共96只个体和一只外群食蟹猴的线粒体DNA控制区576bp的DNA序列，基于这些序列，运用距离法对中国恒河猴的分子进行和遗传多样性进行了分析，我们的研究结果显示，云南、四川和湖南猕猴群体与其它群体存在显著分析，海南群体内遗传多样性最低、四川、广西、浙江、福建和越南群体内遗传多样性较丰富。中国猕猴的分化可能存在三条路线。中国猕猴的遗传多样性较丰富。

Color preferences in gorillas (Gorilla gorilla gorilla) and chimpanzees (Pan troglodytes)

Color plays an important biological role in the lives of many animals, with some species exhibiting preferences for certain colors over others. This study explored the color preferences of two species of ape, which, like humans, possess trichromatic color vision. Six western lowland gorillas, and six chimpanzees, housed in Belfast Zoological Gardens, were exposed to three stimuli (cloths, boxes, sheets of acetate) in red, blue, and green. Six stimuli of the same nature, in each of the three colors, were provided to both species for 5 days per stimulus. The amount of interest that the animals showed toward each stimulus of each color was recorded for 1 hr. Results showed that the apes, both when analyzed as two separate groups, and when assessed collectively, showed significant color preferences, paying significantly less attention to the red-, than to the blue- or green-colored stimuli. The animals' interest in the blue- and green-colored stimuli did not differ significantly. Overall, the findings suggest that gorillas and chimpanzees, our closest living relatives, may harbor color preferences comparable to those of humans and other species. © 2008 American Psychological Association.


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Feature Versus Spatial Strategies by Orangutans (Pongo abelii) and Human Children (Homo sapiens) in a Cross-Dimensional Task

Despite the fact that photographic stimuli are used across experimental contexts with both human and nonhuman subjects, the nature of individuals' perceptions of these stimuli is still not well understood. In the present experiments, we tested whether three orangutans and 36 human children could use photographic information presented on a computer screen to solve a perceptually corresponding problem in the physical domain. Furthermore, we tested the cues that aided in this process by pitting featural information against spatial position in a series of probe trials. We found that many of the children and one orangutan were successfully able to use the information cross-dimensionally; however, the other two orangutans and almost a quarter of the children failed to acquire the task. Species differences emerged with respect to ease of task acquisition. More striking, however, were the differences in cues that participants used to solve the task: Whereas the orangutan used a spatial strategy, the majority of children used a feature one. Possible reasons for these differences are discussed from both evolutionary and developmental perspectives. The novel results found here underscore the need for further testing in this area to design appropriate experimental paradigms in future comparative research settings.

Swimming and diving behavior in apes (Pan troglodytes and Pongo pygmaeus): first documented report

Extant hominoids, including humans, are well known for their inability to swim instinctively. We report swimming and diving in two captive apes using visual observation and video recording. One common chimpanzee and one orangutan swam repeatedly at the water surface over a distance of 2-6 m; both individuals submerged repeatedly. We show that apes are able to overcome their negative buoyancy by deliberate swimming, using movements which deviate from the doggy-paddle pattern observed in other primates. We suggest that apes' poor swimming ability is due to behavioral, anatomical, and neuromotor changes related to an adaptation to arboreal life in their early phylogeny. This strong adaptive focus on arboreal life led to decreased opportunities to interact with water bodies and consequently to a reduction of selective pressure to maintain innate swimming behavior. As the doggy paddle is associated with quadrupedal walking, a deviation from terrestrial locomotion might have interfered with the fixed rhythmic action patterns responsible for innate swimming.

Mirror self-recognition in a gorilla (gorilla gorilla gorilla)

Psychologists have studied self-recognition in human infants as an indication of self-knowledge (Amsterdam, 1972) and the development of abstract thought processes. Gallup (1970) modified the mark test used in human infant work to examine if nonhuman primates showed similar evidence of mirror self-recognition. Chimpanzees (Pan troglodytes) and orangutans (Pongo pygmnaeus) pass the mirror self-recognition test with limited mirror training or exposure. Other species of primates, such as gorillas and monkeys, have not passed the mirror test, despite extensive mirror exposure and training (Gallup, 1979). This project examined a gorilla (G. gorilla gorilla) named Otto in the traditional mark test. Using the modified mark-test, there were more incidents of touching the marked area while Otto was in front of the mirror than when he was not in front of the mirror. These results indicated that Otto was able to show some evidence of selfawareness.

The Influence of Fallback Foods on Great Ape Tooth Enamel

Lucas and colleagues recently proposed a model based on fracture and deformation concepts to describe how mammalian tooth enamel may be adapted to the mechanical demands of diet (Lucas et al.: Bioessays 30[2008] 374-385). Here we review the applicability of that model by examining existing data on the food mechanical properties and enamel morphology of great apes (Pan, Pongo, and Gorilla). Particular attention is paid to whether the consumption of fallback foods is likely to play a key role in influencing great ape enamel morphology. Our results suggest that this is indeed the case. We also consider the implications of this conclusion on the evolution of the dentition of extinct hominins.

Dated co-occurrence of Homo erectus and Gigantopithecus from Tham Khuyen Cave, Vietnam.

Tham Khuyen Cave (Lang Son Province, northern Vietnam) is one of the more significant sites to yield fossil vertebrates in east Asia. During the mid-1960s, excavation in a suite of deposits produced important hominoid dental remains of middle Pleistocene age. We undertake more rigorous analyses of these sediments to understand the fluvial dynamics of Pleistocene cave infilling as they determine how skeletal elements accumulate within Tham Khuyen and other east Asian sites. Uranium/thorium series analysis of speleothems brackets the Pleistocene chronology for breaching, infilling, and exhuming the regional paleokarst. Clast analysis indicates sedimentary constituents, including hominoid teeth and cranial fragments accumulated from very short distances and under low fluvial energy. Electron spin resonance analysis of vertebrate tooth enamel and sediments shows that the main fossil-bearing suite (S1-S3) was deposited about 475 thousand years ago. Among the hominoid teeth excavated from S1-S3, some represent Homo erectus and Gigantopithecus blacki. Criteria are defined to differentiate these teeth from more numerous Pongo pygmaeus elements. The dated co-occurrence of Homo erectus and Gigantopithecus blacki at Tham Khuyen helps to establish the long co-existence of these two species throughout east Asia during the Early and Middle Pleistocene.